Botanica Marina Vol. 45, 2002, pp.

231–242

© 2002 by Walter de Gruyter · Berlin · New York

A Revision of Amansia glomerata C. Agardh, Amansia rhodantha (Harvey) J. Agardh and Melanamansia glomerata (C. Agardh) R. E. Norris (Rhodophyta: Rhodomelaceae)
A. D. R. N’Yeurt
Jeune Equipe Terre Océan, Université de Polynésie Française, BP 6570, Faa’a Aéroport, Tahiti 98702, French Polynesia, nyeurt@upf.pf

Species of Amansia and Melanamansia from the tropical Pacific region have been re-examined, and compared with material from the Indian Ocean, Japan, Taiwan and the Philippines. All specimens examined from the Pacific region (except Hawaii and some from New Caledonia and Fiji) lack pseudo-pericentral cells, thus placing them in Amansia. Melanamansia glomerata is only reported from Hawaii, New Caledonia, Fiji, the Philippines,Vietnam, Malaysia and for the first time in the Indian Ocean from Kenya.Type material of Amansia rhodantha (Harvey) J. Agardh from Mauritius was compared and found not significantly different from the Pacific Amansia material (including type and isotype material of Amansia paloloensis South et Skelton from Samoa, which is reduced to synonymy), thus extending the distribution range of A. rhodantha as a pantropical entity.

Introduction
The genus Amansia was erected by Lamouroux (1809: 322) and is placed in the tribe Amansieae of the family Rhodomelaceae (Hommersand 1963: 335). The tribe is characterised by dorsi-ventral, erect thalli arising in tufts from a common basal disc. Most genera have five pericentral cells around the axial cell (Figs 1, 2) except Kuetzingia Sonder and Protokuetzingia Falkenberg, which have six pericentral cells (Kylin 1956). The two first-formed pericentral cells lie on the dorsal side of the characteristically inrolled apex of the mature lanceolate blades. In Amansia the dorsal and dorsi-lateral pericentral cells further divide to form immediately behind the apex a distromatic wing or ala, whose cells are arranged in alternate layers, with secondary lateral pit-connections (Fritsch 1945: 570). Recently, Norris (1988b, 1995) re-examined species of Amansia in the Indian Ocean and Hawaii, and discovered that some members of this genus differed consistently by having the first and second pericentral cells each producing a lateral pseudo-pericentral cell lying adjacent, but not connected to, the axial cell (Norris 1988b: 214; Fig. 3). Species previously ascribed to Amansia that had these pseudo-pericentral cells were transferred to a new genus (Melanamansia R. E. Norris 1988b: 217). Notably, Hawaiian plants previously ascribed to the common tropical species Amansia glomerata C. Agardh (whose type locality is Hawaii) were transferred to Melanamansia glomerata (C. Agardh) R. E. Norris, while Indian Ocean ‘Amansia glomerata’ were put under the earlier name Amansia rhodantha (Harvey) J. Agardh based

on their lack of pseudo-pericentral cells (Norris 1995). The genus Amansia currently consists of four species, the Caribbean A. multifida Lamouroux (the type of the generic name), A. loriformis R. E. Norris and A. rhodantha (Harvey) J. Agardh from the Indian Ocean and Malaysia, and A. paloloensis South et Skelton from Samoa. Melanamansia currently consists of twelve species, M. daemelii (Sonder) R. E. Norris, M. glomerata (C. Agardh) R. E. Norris, M. japonica (Holmes) R. E. Norris, M. mamillaris (Lamouroux ex C. Agardh) R. E. Norris, M. pinnatifida (Harvey) R. E. Norris, M. pumila (Sonder) R. E. Norris, M. scalpellata (Tanaka) R. E. Norris, M. seagriefii R. E. Norris, M. serrata (Harvey) R. E. Norris and M. mitsuii (Tanaka) Yoshida. Following these revisions by R.E.Norris,several authors (Yoshida et al. 1995,N’Yeurt et al. 1996,Payri and N’Yeurt 1997, Millar 1999, Millar et al. 1999) made the nomenclatural change from Amansia glomerata to Melanamansia glomerata in their checklists and herbarium collections from the tropical Pacific, but until recently no detailed studies verifying these changes as correct have been done. In 1995, samples of ‘Amansia glomerata’ from Rotuma Island (north of Fiji) were sent to Dr R.E.Norris,who confirmed (pers. com.) that pseudo-pericentral cells were present, and on the basis of this assumed finding the name Melanamansia glomerata was used in later publications (N’Yeurt 1996,N’Yeurt et al. 1996).Later,in the course of on-going investigations on the Samoan algal flora, South and Skelton (1999) found a species of Amansia lacking pseudo-pericentral cells, which they described as new (A. paloloensis South et Skelton), while in Malaysia and Vietnam (Masuda et al. 2000, Masuda

Sakai and S. Payri (8/84. M. Skelton. P. 3892. 1997. 3896. 2. Prepared slides were examined using a Zeiss compound microscope. Amansia rhodantha (Beqa. leg. a = axial cell. deep algal rim. 25 October 1984 (SAP 046978). 5481L (spermatangial)]. The results of these investigations are described below. 19 Mar. 3895. 5480L (cystocarpic). Punaauia.‘Al’ and ‘CG’ refer to specimens housed at IRD. and a camera-lucida attachment was used to make anatomical drawings. P. 13 May 1982. snorkelling or SCUBA diving. – French Polynesia: Moorea. 1998. Baba (SAP 056160). . – Samoa: Palolo Deep Reserve. leg. SUVA-A 2402 (Lectotype) and liquid-preserved isotypes 5479L (tetrasporic). Tahiti. 2 Jul. Nouméa.) recent studies found both Melanamansia glomerata and Amansia rhodantha (Harvey) J. R. M. leg. and stored in 5% buffered Formalin in seawater prior to shipment to the laboratory. Enomoto (SAP). New Caledonia. leg. Arrangement of pericentral cells in species of Amansia and Melanamansia (modified from Norris 1988b. a detailed morphological and taxonomic study on some selected members of the Amansieae (Wilson and Kraft 2000) echoed the opinion that the genus Amansia in Australia was in need of critical revision. leg. com. Okinoerabujima Island (60 km north of Okinawa) 27 Sept. 6 Jun. Fig. P. leg. M. 1989. Fig. Skelton. 1998. N’Yeurt (SUVA-A 5128). A. Melanamansia glomerata. in EPHE). Kagoshima: 9 August 1959. Furusato. A. Amansia rhodantha and Amansia glomerata.Agardh to co-exist in the flora. Rintels (SUVA-A 3891. Baba (SAP 056163). 1998. Tanegashima Island. 3. 1998. (1981). Fig. N’Yeurt (UPF 584). SUVA-A 5436L). Skelton and Dave (SUVA-A 1567). Macrophotography was done with a Nikon F2A in conjunction with Kodak Plus-X pan 125 film. 1998. Skelton and Dave [SUVA-A 1565 (holotype). Rarotonga. U. E. otherwise as in Holmgren et al. 18 Sep. – Cook Islands: Ngatangiia. 1–5 = order of formation of pericentral cells.. 3898. C. developed and printed in the laboratory. A. Figs 1–3. A. Palolo Deep. Faletose (SUVA-A 3894. leg. Material and Methods Specimens were collected by reef-walking. SUVA-A 1566 (isotype). leg. 1984. Okidomari. Most recently. Okinoerabujima Island (60 km north of Okinawa). A. 11 Jul. it was decided to revise records in the light of recent findings. D. leg. showing intrusion of the first derivative of the first-formed pericentral cell into the axial ring (Suva. Tombleson and T. 2 Jul. N’Yeurt (UPF 347). Taharaa. – Taiwan: Anon. pers. 3897). and stained with cotton blue in lactophenol or 1% acidified aniline blue in 60% clear corn syrup. in particular paying attention to important anatomical features such as the presence or absence of pseudo-pericentral cells and the number of segments between lateral veins of the blade (Norris 1995). 3899). 3893. A. N’Yeurt In order to clarify the taxonomy of species hitherto ascribed to Melanamansia glomerata. Afiti and D. A. Y. leg. Photomicrographs were obtained using an Olympus BX-50 compound microscope. Fiji. Amansia rhodantha. Results Representative material examined Amansia rhodantha – Japan: Hanasado Beach. Tahiti. 9 Feb. Fiji. 1999. ps = pseudo-pericentral cells). cross section of the lectotype from the Hawaiian Islands (LD 42600) showing the first two dorsal pericentral cells each developing a pseudo-pericentral cell (ps) that lies adjacent to the axial cell. P. leg.232 A. Sections were made using a freezing microtome. 1. SUVA-A 5438L).

Salayer.Negros Oriental. Eagles Nest. CG 77b). Yasawa Group. Magruder (US 086459). 1997. 1996: 472. – Australia: Heron Island QLD. 1067). Garrigue (CH1). – Mozambique: Ponta Abril. Average values of selected characters for Amansia paloloensis. leg. (MURU. South (SUVA-A 5544. 1819.Moreland (US 50851).Negros Oriental.d. leg. leg. – Fiji: ‘Feejee’. Cannon (SAP 052062). 1988. leg. Garrigue (CG 77c. SIPHILEXP-78 (US 088531). 2000: 188. L 0108598). Potipot Is. – Philippines:Gao-oa.. J. 14 Feb. Hapmak. Huisman (MURU. R. Kasahara (SUVA-A 1012. leg. Basay. 5 June 1967. Agardh). 07 Oct. R.9 11. 84. leg. Oahu. 1995. Villeneuve (SUVA-A 1737).leg. leg.Calindagan Reef. A. leg. S. 1994 (SUVA-A 797). Catanduanes.P. JH240). Vatualailai. 1980. Kauai. Wilkes Exploring Expedition (US 04057b) (doubtful record. 18 March 1992. leg. – Kenya: Mombassa. 13 July 1972.d.W. leg. 1834).7 54.23 May 1978. Suva Reef. leg. 29 May 1978. leg. South in lit. Lawai Kai. Cordero and Tuangpalan (US 091394). Jul. leg. Tantabiddi. leg. H. leg. leg.2 53. Inhaca Peninsula. leg.leg. 1985. Huisman (CWC 1997. SIPHILEXP-79 (US 088533). leg. Garrigue (SUVAA 5436L). R. W. Isaac (L 0108595). 9 Sept. 25 September 1999. Banilad.. 1995. LD 42600 / 0772). Ballou (SUVA-A 5438L). Hildebrandt (L 0108600). Tangangge. Candelaria Zambales. Negros Oriental. W. 1979. 21/22-Sept. J. A. leg.d. 1983. A. – New Caledonia: Lajeu Sudouest. 6 Jun 1978. SIPHILEXP-78 (US 088530).9 18. Taxonomic results Amansia rhodantha (Harvey) J. Daida (US 086755). A. Lopta. A. Barrow Island. 1069). Kasahara (SUVA-A 1013. 10 July 1992. Plongeurs IRD (AL 288a. Oahu.Dumaguete City. Table I presents the average results of morphological measurements carried out on Amansia rhodantha. Unia. – Philippines: Actin. 3 Feb.Canipo Is. leg. Littler (US 163490). W. 27 Dec. Species Thallus height (cm) Blade width (mm) Blade length (mm) Serrations (1 –3)* Cortication (1 –3)* Rosettes (1 –3)* Amansia rhodantha Amansia paloloensis Melanamansia glomerata 39. Nanuyalevu. leg. leg. P.Siaton. Cordero and Tuangpalan (US 091395). South (SUVA-A 2460). Kipukai. Magnesia Virae. Melanamansia glomerata – Hawaiian Islands: Hawaiian Islands. 1984. 1838–42. C. M. P.Revision of some Amansia and Melanamansia species 233 Table I. 24 Nov. 1981. J. anon. 1995. 347). 84. A. (US 02890). A. Beqa Lagoon. Agardh 1841: 26. SIPHILEXP-78 (US 013760). Viti Levu. – Mauritius: Delesseria rhodantha Harvey (holotype) (= Amansia rhodantha): Cap Malheureux. 26 Nov. J. 1838–42. – Kenya: Mombassa.. Barrow Island. Dumaguete. S. Snellius-II (L 0108602). B79). E. Bahati. Lectotype. Montebello Islands. W. 21 Sep. leg. Amansia rhodantha: n. leg. Oahu. leg. Huisman (MURU. leg. Hakula. E. C. 1954. Isaac (L 0108594).14 May 1979. 16 March 1991. J. Huisman. Yasawa Islands. leg. n. 14 March 1998. 1985. Mauritius. Naukathura Island.. R. Huisman (MURU. 17 Mar.. M de Robillard (L 0108599). G. Hildebrandt (L 0108596. Chesterfield. 28 Nov. leg. Gaudichaud (‘Amansia glomerata’. Viti Levu. Cribb (SAP 051714). Villeneuve (SUVA-A 1774). – Lord Howe Island: 18 Aug. Jurien Bay WA. 10 May 1978. 25 Feb. N’Yeurt (SUVA-A 346.Malo. M. 1980. leg. – New Caledonia: Lajeu Sudouest. leg. see text). H. Bay of Sanggar. E. 1973.Ilocos Norte. NR 248).Siquijor Is.1 22. Viti Levu. 16 Nov. leg. figs 1–11 (as Amansia glomerata C. Rukuruku Village. 1982. leg. c. 1876. leg. figs 37–42. 24 April 1996. 3 and 15 Oct. Carlson (SUVA-A 281). leg. B.5 3. A. 1965. leg. G. Wilkes Exploring Expedition (US 04057a).d.. n.11 Feb.. D. fide G. leg. R127). G. Silva et al. Namada. leg. 1876. Sumbawa. Masuda et al. J. B262). 1982. J. leg.Solong-On. Jul. leg. paloloensis and Melanamansia glomerata specimens examined in this study. leg. 84 (L 0108603).Pagodpud. 17 Jul. – Indonesia: Cape Batu Kerapo. Keats. leg. South (SUVA-A 5431). Richer de Forges (AL 288d-CG 77a). Hanauma Bay Beach Park. R. Ovalau. 16 Oct. Norris 1988b: 211. Diani Beach. 10 Aug. Amansia rhodantha and Melanamansia glomerata surveyed in this study. SIPHILEXP-78 (US 088534). AL 288b). H. in TCD (Telfair.). 3 Jun 1978. C. C. Snellius-II (L 0108601). leg. M. leg. 1979. – Fiji: ‘Feejee’. M. leg. Skelton and G. Negros Oriental. Ningaloo. N’Yeurt (SUVA-A 5437L). 1984. 06 Sept. 3 and 5 March 1992. Ti Tree.. n. Farlow (L 0108597). 30 Jun.Cuyo.6 3. leg. leg. Kauai. Suva Barrier Reef. n.Palawan. B. B.7 2 2 1 2 2 2 2 2 2 * As measured on an index of (1 = little) to (3 = pronounced) – Rotuma: Hoféa.3 2. P. 15 Oct. SIPHILEXP-78 (US 088532). G. Figs 4–10 .d.

Fig. Figs 4. Type material of Delesseria rhodantha Harvey. Detail of tetrasporangial stichidia on lateral endogenous branch. Fig. and a pair of budding branchlets from the midrib in the lower half. Axial structure of blade.5 mm). Secondary branchlet. SUVA-A 5437L. Surface view of segment cells of blade alae. Habit of mature plant from the type locality. showing both lax and dense growth forms. D. Rehydrated type material of Delesseria rhodantha Harvey. Type material of Delesseria rhodantha Harvey. N’Yeurt Figs 4–10. 6. Fig. . 7. from Rotuma Island. 5. 8. showing marginal serrations. Indian Ocean. in TCD (scale bar = 25 µm). 9. in TCD (scale bar = 2. South Pacific (scale bar = 10 µm). Mauritius.234 A. US 02890 (scale bars = 3 mm). R. 10. Amansia rhodantha. showing axial cell (a) surrounded by five pericentral cells. in TCD (scale bar = 200 µm). Fig.

Fig. 16. Note marginal cystocarps (arrowheads). Segment cells of blade alae in surface view. SUVA-A 3892 (scale bar = 2. SUVA-A 1565 (scale bar = 25 µm). showing lateral endogenous branches and midrib. SUVA-A 3892 (scale bar = 12 mm). Compare with Fig. 14. with numerous rosette axes. Fiji. 12. Fig. SUVA-A 5479L (scale bar = 1 mm). Tetrasporangial stichidia of Amansia rhodantha from Beqa. .Revision of some Amansia and Melanamansia species 235 Figs 11–16. Holotype of Amansia paloloensis. showing both lax branching and the presence of rosettes (arrowheads) (scale bar = 4 mm). Fig. Fig. Fig. Detail of morphologically distinct mature rosette and corticated axis. Detail of blades. Habit of mature plant. Detail of the holotype (SUVA-A 1565). 11. Amansia paloloensis from Samoa. Fig. 15. SUVA-A 5436L (scale bar = 50 µm).5 mm). 13. 4.

Kenya. Silva et al. 2. Plant from Mombassa. pinnately arranged marginal stichidia up to 340 µm long and 290 µm in diameter. pl. Agardh) R. Fig. 197. Tetrasporangia occur in flattened. The number of segments between alternating veins (determinate branches covered by the corticating pericentral cell derivatives. Fig. they are up to 4 cells long and non-vesiculate. 118C–J. A distinct midrib up to 1 mm wide is present in the lower half of ultimate branchlets. Kylin 1956: 544. holotype in TCD. 436B. with a range of 3–6. – Melanamansia glomerata (C. figs 2–14 (type locality: Palolo Deep Marine Reserve. E. Norris 1995: 66) is on average 4. Coppejans and Millar 2000: 333 (unverified record). N’Yeurt Basionym: Delesseria rhodantha Harvey 1834: 151–152. US 4057b (scale bar = 1 mm). 6. containing up to 10 Figs 17–24. and are composed of lanceolate to ovate blades to 100 mm long and 3 mm wide. Fiji. Marginal teeth are up to 1 mm long and 445 mm broad. 18. Axial structure of plant from Lajeu. South-East Asia. fig. Isaac 1956: 188. figs 14–29. Okamura 1900: 71. Figs 11–16 Misapplied names: Amansia glomerata C. Dublin). to 0. with in-rolled leaf apex. Payri and N’Yeurt 1997: 897. Weber-van Bosse 1923: 369. Agardh 1824: 194. in up to 10 regularly disposed pairs. New Caledonia. denuded below. up to 750 µm long and 250 µm broad. 1995: 150. Note that the stichidia is twice laterally branched. 1. figs 1–13. US 091395 (scale bar = 5 mm). The central axial cell is 28–30 µm in diameter. Cribb 1983: 106. Fig. Amansia paloloensis South et Skelton 1999: 247. with more or less pronounced marginal serrations 1–4 mm long (which represent endogenous branches) always present. with a central axial cell 28–30 µm in diameter surrounded by five pericentral cells 36–40 µm in diameter (Figs 1. Blade margins are smooth. R. 1987: 61. Payri and Meinesz 1985: 511. Blade of plant from Lajeu. Procarps and curved spermatangial heads are 200–450 µm in diameter and are dorsal on the involucral tips of endogenous lateral branches. becoming narrower and disappearing towards the apex. Fig. with a range of 1–5. Agardh 1824: 247. Yoshida et al. and are crisp and lanceolate. Falkenberg 1901: 416. composed of two overlapping layers of elongate hexagonal cells in transverse rows. Ultimate branchlets are ecorticate. N’Yeurt et al. Agardh) R. Millar et al. pl. and two displaced rows of regularly arranged hexagonal alar cells 20–29 × 92–129 µm in surface view. Tsuda and Wray 1977: 103. 1999: 573. Norris 1995: 67. 20. fig. figs 19–21. 0. Branching is lax to profuse and usually forms deep-red leafy rosettes of secondary branchlets when fully mature. Plant from Potipot Island. and the apex is characteristically in-rolled. AL288b (scale bar = 5 mm). Mauritius. Isaac and Isaac 1968: 24. 19. The stem of plants is thick and cartilaginous. New Caledonia. New Caledonia. Garrigue and Tsuda 1988: 63. Synonymy: Rytiphlaea rhodantha (Harvey) Decaisne 1842: 358. Philippines. showing smooth margins and elongate shape. Blades of plant assumed to be from the Fiji Islands (see text). Norris 1995: 67. figs 1–3 (type locality: Cap Malheureux. while cystocarps 700–1000 µm in diameter occur usually singly and terminally on endogenous branches. Jaasund 1976: 131. Deciduous. CG77a (scale bar = 1 mm). fig. 6E (unverified record). Detail of blade of plant from Mombassa. plate CXXVI. Internal structure is cellular. Tropical Pacific Ocean. up to 35 mm long and 6 mm broad. 1. Trichoblasts are sparse. Japan. Apia. Fritsch 1945: 570. 128. Individual tetrasporangia are spherical and tetrahedrally divided.8 mm in diameter. East Africa. D. L 0108598 (scale bar = 5 mm). Melanamansia glomerata (C. SUVA-A 5431 (scale bar = 50 µm). figs 1–11. surrounded by five pericentral cells. 19). 267. Fig. Vietnam. 21. 77–110 µm thick.4. 198. Kenya.5 µm in diameter that lies adjacent to the axial cell (Figs 3. each cell is about 20–20 × 87–100 µm in surface view. 20. 100–112 µm in diameter. Plants are brownish-red. Norris 1995: 66) is usually 4. 185. 9). 25–42 × 83–125 µm. The main axis is irregularly branched. Plant from Unia. The number of segments between alternating veins (determinate branches covered by the corticating pericentral cell derivatives. 31 fig. Philippines. A central midrib is present. Fig. E. Melanamansia glomerata. with the first two dorsal pericentral cells each developing a smaller pseudo-pericentral cell 13. 22. New Caledonia. Norris 1988b: 211.8–14. Samoa). e–i. fig. 85–110 µm thick. colourless and uniseriate dorsal trichoblasts up to 8 cells long develop on every second or third axial cell (or on every segment in determinate marginal branchlets). pl. Womersley and Bailey 1970: 336. 1996: 87. Distribution: Tropical Indian Ocean. The thallus is rose-red. N’Yeurt 1996: 428. showing pseudo-pericentral cells (arrowheads) surrounding (but not directly connected to) axial cell (a). Kenya. up to 100 (on average 40) mm high. and are initially vesiculate. Secondary blades develop as successive pairs on the dorsal side of the midrib. .236 A. Tetrasporangial stichidia of plant from Oahu. 17.5–0. stem-like and rigid in basal portions. on average 53 mm high. Figs 17–24 Basionym: Amansia glomerata C. or with regular serrations (endogenous branches). Yamada and Tanaka 1938: 86. 212a. 4B. Known distribution: Hawaiian Islands. 7 mm broad. CG77a (scale bar = 10 µm). L 0108598 (scale bar = 700 µm). Millar 1999: 523 (unverified record). Carposporangia are lanceolate. Abbott 1999: 404. pl. fig. XXV. Lewis and Norris 1987: 22. pl. figs 127. 23. Fig. Tetrasporangial stichidia are pinnately arranged on either side of blade margins. Fig. 24. fig.

procarps and cystocarps occur on endogenous marginal branches. In all cases. and stained the herbarium mounting paper. Spermatangial heads are curved and arranged in marginal clusters. .Revision of some Amansia and Melanamansia species 237 pairs of tetrahedrally-divided tetrasporangia which are 50–100 µm in diameter. the colour of the Melanamansia plants was noticeably brownish-red.

having tight rosettes as well as very lax. length of the blades. a single record for this genus exists (collected from ‘Feejee’ by the Charles Wilkes U. Kenya. and found a significant number of thalli with morphologically distinct primary corticated axes and relatively tight and lax rosettes of secondary blades on rosette axes (Figs 11. from Melanamansia glomerata was found: when preserved in 4% Formalin-seawater and exposed to ambient light for several days. Eastern Australia.13). rhodantha (South and Skelton 1999: 249). 3) which distinguishes that plant from A. when pseudo-pericentral cells were found to be absent) are considered conspecific with Amansia rhodantha. From this study. an additional field test to distinguish Amansia spp.). more coriaceous branchlets with less apparent midribs (this study). rhodantha. while the other (US 04057b) is Melanamansia glomerata. N’Yeurt Discussion The holotype of Delesseria rhodantha Harvey An examination by the author of a rehydrated blade fragment of the holotype of Delesseria rhodantha Harvey in TCD (Fig. it appears to be conspecific with the latter species and thus extends the status of A. paloloensis. and a weakly developed midrib as characters distinguishing A. paloloensis showed no major difference in the range of characteristics among them. the Philippines. paloloensis. Rotuma. generally smaller. paloloensis from the type locality. including the presence of both tight and lax rosettes of secondary branches and alar cells on the primary axes. The drawing of the holotype (South and Skelton 1999: 246. rhodantha from the Indian Ocean and other localities. Smith (1979: 38–40) points out that the accounting and . On the other hand. paloloensis from A. as well as Amansia species from Taiwan. and these authors considered the lack of highly corticated axes. Mozambique and Western Australia. rhodantha from the type collection and localities in Mauritius. Kuetzingia. which is welldocumented and illustrated in the protologue of the species. a comparison of material of A. However. Exploring Expedition. apparently because the brown pigments (presumably polyphenols) are less susceptible to the bleaching action of the preservative (as is the case in species of brown algae such as Sargassum and Turbinaria). a re-examination of the lectotype of Amansia glomerata C. since the presence of pseudo-pericentral cells is an inconsistent attribute in species of Neurymenia. Vidalia. Melanamansia glomerata in the tropical Pacific Wilson and Kraft (2000: 366) commented that the generic segregation of Melanamansia from Amansia is probably not warranted.S. The status of Amansia paloloensis South et Skelton South and Skelton (1999) described a new species of Amansia from Samoa. The Wilkes collection consists of two plants. R. the Cook Islands and French Polynesia (Table I) with A. is comparable to A. Japan. plate CXXVI. serrate branches and lax ‘rosettes’ of secondary blades issued from the weakly developed midrib. while Melanamansia becomes dark brown. loriform blades reminiscent of some specimens of A. 12). and a cross section of the blade clearly shows the five pericentral cells around the axial ring. More importantly. 7) shows features similar to those seen in blades of South Pacific material previously ascribed to Amansia glomerata. These plants also had blades with well-developed midribs in the lower half (Fig. indicating that the morphology of the species is influenced perhaps by environmental factors. New Caledonia. rhodantha from Western Australia (MURU B262. fig. one of which (US 04057a) is Amansia rhodantha. M. regularly alternate marginal serrations which are not as pronounced as in Amansia rhodantha and which are often absent. material from the Tropical Pacific and Indian Oceans previously known as Amansia glomerata (or misidentified as Melanamansia glomerata. Masuda pers. additional distinguishing characteristics of Melanamansia are a darkly coloured brown thallus which stains the mounting paper when dried (possibly linked to differences in chemical composition. rhodantha as described and illustrated by Norris (1988b: 211). On the basis of these observations. D. Fig. In the Fiji Islands. Characters such as the degree of rosette formation. com. figs 1–4) indicate features quite similar to those found in the Amansia species considered in this study from localities in both the Indian Ocean and the tropical Pacific Ocean. between 1838 and 1842 (US 04057b. Agardh from Hawaii (LD 42600 / 0772) clearly shows the presence of paired dorsal pseudo-pericentral cells in a blade cross section (Fig. The present author has re-examined the holotype.238 A. and absence of pseudopericentral cells. 23) but the plant has not been collected in Fiji waters since. 14) and corticated rosette axes (Fig. and trichoblasts lacking protective vesicles when young (Norris 1988b). While there is no doubt as to the generic identity of the latter specimen. degree of cortication of the midrib and position of the reproductive stichidia are equally quite variable in plants from the same and similar populations. rhodantha as a pan-tropical entity. thicker. 2) indeed shows a plant with loriform. Now that the Samoan species has been compared with genuine herbarium material of A. The reproductive and trichoblast anatomy of A. Amansia tends to bleach to translucent white. The habit and description of the holotype by Harvey (1834: 151. Fiji. and Ceramium. with for instance A. the absence of blades organized in morphologically distinct tight axes around the parent axis. isotype and other collections of A. MURU 248).

com. However. when present). in some cases. Masuda. Indian Ocean Amansia rhodantha also lacked such cells. 13). pers. Payri and N’Yeurt 1997) were reexamined. alternate marginal proliferations (which represent non-determinate or endogenous secondary branchlets) 2–5 mm long (Fig. the presence or absence of bladelets in rosettes. and is backed by other morphological and biochemical differences (Norris 1988b. in the case of A.g. so it cannot be confirmed at this stage if M. and would seem to function as attachment haptera. The presence or absence of these pseudo-pericentral cells hence appears to be a good generic criterion. while in Malaysia Masuda et al. Also. fig. From an examination of the specimens at hand. 2). and found to lack pseudo-pericentral cells. and in some instances lacking altogether. it was observed that rosettes of blades along the secondary axes. type of endogenous branching. older plants tend to have thicker. more corticated secondary and primary axes with tight rosettes. The illustration by Harvey of the holotype of Delesseria rhodantha (1834. Herbarium specimens of ‘Amansia glomerata’ from southern Japan housed in SAP (Hokkaido University) were examined. Norris (1988b: 211) reported that rosette axes in .). 5. Herbarium and liquid-preserved samples of Amansia and Melanamansia species from the Pacific. the description of the species states that branches have the same shape as the main axis (i. irregularly dichotomous proliferations lack alar cells. When pseudo-pericentral cells were present. which was the main reason for separating the species from A. M. the Philippines. both have an ala) and are arranged in ‘lax’ rosettes (South and Skelton 1999: 247. fig. It would seem that perhaps shallow. These ventral. more exposed conditions (such as on fringing reefs) favour tight rosette formation (which is more hydrodynamically resistant) while more lax arrangements are possible in deeper. pseudo-pericentral cells were universally absent. Validity of current criteria to distinguish the Amansieae Characters that have proved useful in distinguishing species in the Amansieae include the presence or absence of pseudo-pericentral cells. and the presence of tight and lax rosettes of secondary branches on the same plant (Figs 11–13). Wilson and Kraft 2000). which is a characteristic feature of both Amansia and Melanamansia (Norris 1988b) can vary from lax to tight. 7). glomerata occurs in the same habitat as Amansia rhodantha (e. paloloensis revealed a wide range in the degree of rosette formations and cortication of the secondary axes and presence of alae on primary axes. 1) shows both the presence of tight and lax rosettes of secondary branches. fig. position of the reproductive axes and the distinctness of the axial ring in rosette axes (Norris 1988a. However. while younger plants have less cortication and more lax or absent rosettes of blades. adding to the confusion between the two genera. Melanamansia glomerata is unlikely to be confused with other Melanamansia species because of its unique combination of characteristics (Norris 1988b. however it is clear that the degree of rosette formation and cortication of the axes is not a reliable taxonomic character at the species level in Amansia. Indeed.Revision of some Amansia and Melanamansia species 239 recording of the Wilkes Expedition collections was not very satisfactory. and some samples from Kenya. an examination of the holotype and liquid-preserved and pressed isotypes of A. rhodantha from Mauritius and Western Australia (compare Figs 4. Ecological studies would be needed to verify these hypotheses. Fiji and New Caledonia (Table I). perhaps an adaptation to the exposed habitats where the plants came from. All Amansia rhodantha specimens examined had fairly prominent. calm waters (such as blue holes. except the Hawaiian samples. M. In Melanamansia glomerata these marginal serrations were much less pronounced (but more regular than in Amansia rhodantha. rhodantha. rhodantha. the vesiculate (in Amansia) or non-vesiculate (in Melanamansia) nature of trichoblasts. For instance. and hence the precise locality and date of this Melanamansia collection from Fiji is unknown. There is a possibility it might have been mixed with collections from neighbouring localities such as the Solomon Islands. they were clearly apparent and about half the size of pericentral cells (Fig. with the blades smooth and superficially reminiscent of those of some Sargassum species (Figs 22–24). 1996. Fijian and French Polynesian specimens previously ascribed to Melanamansia (N’Yeurt et al. which also had a consistently thicker thallus. The author has unsuccessfully tried to obtain material from the latter locality. in the case of Amansia paloloensis). in New Caledonia and Kenya). (2000: 188. and a main axis with alar cells not very distinct from secondary branches. 1995).e. 19). Generally. degree of marginal serrations and thallus flattening. both on the same plant and within individuals in similar populations. In most instances. pl. paloloensis. and found to consistently lack pseudo-pericentral cells and a similar observation was made on samples from Taiwan. environmental factors seem to play a role in determining the degree of rosette formation. thus placing them in Amansia. 41) similarly report the absence of pseudo-pericentral cells in A. L 0108603). CXXVI. some specimens were virtually indistinguishable from specimens of A. Indonesian and Indian Ocean region were sectioned and examined for pseudo-pericentral cells. glomerata occurs in the Solomon Islands. Proliferations from the ventral midrib (aside from secondary branchlets) were only found to be abundant in some of the Indonesian Amansia rhodantha specimens from Salayer (e.g.

16) and Melanamansia show no particular difference. in elongate flattened and pinnate stichidia which are usually on lateral marginal branchlets. D. Norris (1995: 66) counted the number of segment cells between these alternating veins for several species of Melanamansia and Amansia. as well as more corticated axes. B. rhodantha specimens observed in this study. but can also occur apically. a number of preliminary conclusions can be made: A. Cover cells usually totally cover the developing sporangia. rhodantha and A. com. and occur in two overlapping and slightly offset layers radiating on either side of the midrib. There appears to be no notable difference in the range of thallus size. However. with most specimens having segment lengths of about 100 µm. At this stage there seems to be no largely significant difference in the number of segments between alternating veins in the specimens of A. as the axial ring was still discernible in all sections made. only being more or less surrounded by cortication. 25.4 (Norris 1995). One feature which was noted in Melanamansia glomerata is that individual tetrasporangial stichidia are often laterally branched once or twice (Fig. Melanamansia glomerata from all localities studied have consistently smoother. These can be easily detected in the unrolled apical portions of the blades by the developing trichoblast terminating each such alternating ‘vein’ (Norris 1995 and pers. Fig. paloloensis. but the range is reportedly less than in Amansia rhodantha.). . which was also recorded by Norris (1988b) for Hawaiian material. Biogeographical map for Amansia rhodantha (circle) and Melanamansia glomerata (triangle). These differences reinforce the conspecificity of the Amansia rhodantha and A. as well as between A. usually 3 or 4 with an average of 3. smaller and narrower blades. N’Yeurt Amansia from Natal have an ill-defined axial row of cells. Hence the distinctness of the axial ring was not found a reliable taxonomic character among the species examined. 15). paloloensis specimens studied. Cells of the blade alae in both Amansia and Melanamansia are regularly elongate-hexagonal. blade dimensions. In A. which are in fact determinate branches covered by the pericentral cell derivatives.240 A. An examination of these elegant cells in species of Amansia (Figs 10. with the length of segments ranging from 80 to 120 µm.3. the number ranged from 3 to 7. it was noted that in Melanamansia the range was more restricted. From Table I. which was not seen in Amansia rhodantha (Figs 6. cortication of axes and rosette formation between specimens of Amansia rhodantha studied. R. Tetrasporangial stichidia are basically similar in both Amansia rhodantha and Melanamansia glomerata. 20). This feature was examined on a range of specimens of Amansia from the South Pacific and the Indian Ocean but no significant difference was seen in these specimens. and found them relatively constant. as well as the generic distinctness of Melanamansia glomerata. This also varied according to the degree of rosette formation and age of the plant. An additional potentially useful character is the number of segments between lateral ‘veins’ on the blades. In Melanamansia it averages 4. rhodantha studied. showing the currently known distribution. in that they are typically rhodomelaceous with pairs of tetrasporangia in up to ten rows.

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